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Gastropoda) of the Islands of Sao Tome and Principe, with New Records and Descriptions of New Taxa

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Gastropoda) of the Islands of Sao Tome and Principe, with New Records and Descriptions of New Taxa This is the submitted version of the article: “Holyoak, D.T., Holyoak, G.A., Lima, R.F. de, Panisi, M. and Sinclair, F. 2020. A checklist of the land Mollusca (Gastropoda) of the islands of Sao Tome and Principe, with new records and descriptions of new taxa. Iberus, 38 (2): 219-319.” This version has not been peer-reviewed and is only being shared to comply with funder requirements. Please do not use it in any form and contact the authors (e.g.: [email protected]) to get access to the accepted version of the article. 1 New species and genera and new island records of land snails (Gastropoda) from the islands of São Tomé and Príncipe Nueva especies de .... David T. HOLYOAK1, Geraldine A. HOLYOAK1, Ricardo F. de LIMA2,3, Martina PANISI2 and Frazer SINCLAIR3,4 Recibidio el ... ABSTRACT Seven species of terrestrial Gastropoda are newly described from the island of São Tomé and six more from the island of Príncipe. The genera involved are Chondrocyclus (Cyclophoridae), Maizania and Thomeomaizania (Maizaniidae), Pseudoveronicella (Veronicellidae), Nothapalus (Achatinidae: subfamily undet.), Gulella and Streptostele (Streptaxidae), Truncatellina (Truncatellinidae), Afroconulus (Euconulidae), Principicochlea gen. nov., Principotrochoidea gen. nov., Thomithapsia gen. nov. and Thomitrochoidea gen. nov. (Urocyclidae). Most of these are from natural forest habitats and are likely to be single- island endemics. Apothapsia gen. nov. (Helicarionidae) is also described to accommodate two previously known species. Additional new island records are of ten species on São Tomé, one on Príncipe alone and two more on both islands. These include six species of "microgastropods" with wider ranges in tropical Africa that are likely to be hitherto overlooked parts of the indigenous fauna and six anthropogenic introductions; Pseudopeas crossei previously known only from Príncipe and Bioko is newly recorded on São Tomé. More detailed taxonomic notes are provided on identification and delimitation of species and genera in Cyclophoridae/ Maizaniidae, Veronicellidae, Achatinidae (for Petriola), Helicarionidae, and on the forgotten but apparently valid Pseudoveronicella thomensis (Veronicellidae). The first comprehensive checklists since the works of Germain in 1916 are presented for the terrestrial Mollusca of both islands, comprising 86 species in total, with 59 known for São Tomé and 45 for Príncipe. RESUMO Sete novas espécies de gastrópodes terrestres são descritas da ilha de São Tomé e seis da ilha do Príncipe. Os géneros envolvidos são Chondrocyclus (Cyclophoridae), Maizania e Thomeomaizania (Maizaniidae), Pseudoveronicella (Veronicellidae), Nothapalus (Achatinidae: subfamília indet.), Gulella and Streptostele (Streptaxidae), Truncatellina (Truncatellinidae), Afroconulus (Euconulidae), Principicochlea gen. nov., Principotrochoidea gen. nov., Thomithapsia gen. nov. e Thomitrochoidea gen. nov. (Urocyclidae). A maioria destes registos foram feitos em habitats de floresta natural e provavelmente são endémicos de uma única ilha. Apothapsia gen. nov. (Helicarionidae) também é descrito para acomodar duas espécies já conhecidas. 10 espécies adicionais são novos registos para São Tomé, uma para o Príncipe e duas para ambas as ilhas. Estes incluem seis espécies de "microgastrópodes" com ampla distribuição na África tropical, que provavelmente são parte da fauna indígena que tinha passado despercebida, e seis introduções antropogénicas; Pseudopeas crossei anteriormente conhecida apenas do Príncipe e Bioko é um novo registo para São Tomé. São fornecidas notas taxonómicas mais detalhadas sobre a identificação e delimitação de espécies e géneros de Cyclophoridae/ Maizaniidae, Veronicellidae, Achatinidae (para o género Petriola), Helicarionidae e para a 2 esquecida, mas aparentemente válida Pseudoveronicella thomensis. São apresentadas as primeiras listas compreensivas de moluscos terrestres para ambas as ilhas desde os trabalhos de Germain em 1916, compreendendo 86 espécies no total, com 59 conhecidas para São Tomé e 45 para o Príncipe. RESUMEN Se describen x especies nuevas de Gastropoda terrestre de la isla de Santo Tomé (to be completed) 1Quinta da Cachopa, Rua da Barcoila no. 274, 6100-014 Cabeçudo, Portugal. Corresponding authors, e-mail: [email protected] 2Centre for Ecology, Evolution and Environmental Changes (Ce3C), Faculdade de Ciências, Universidade de Lisboa, 1749-016 Lisboa, Portugal. 3Departamento de Biologia Animal, Faculdade de Ciências, Universidade de Lisboa, 1749- 016 Lisboa, Portugal. 4Fauna & Flora International, Cambridge, U.K. 5Fundação Príncipe, Santo António, Príncipe Island, Republic of São Tomé and Príncipe. __________________________________________________________________________ 3 INTRODUCTION The Republic of São Tomé and Príncipe comprises two mountainous islands lying close to the Equator in the Gulf of Guinea, off the west coast of central Africa. Both islands form part of the Cameroon Line of volcanos, and both arose due to volcanic activity during the Tertiary. They rise from oceanic depths and have always been isolated from the nearest African coasts, which are no closer than 220 km and 255 km distant at the present day. Their natural vegetation was tropical rainforest, although cultivation has replaced most of it on the lower ground since human colonisation began in the fifteenth century (JONES & TYE, 2006). Both islands support rich land-snail faunas with a high proportion of endemic species. GASCOIGNE (1994a, 1994b) reported the faunas as comprising 39 species for São Tomé and 32 species for Príncipe, although he never published a full list for either island. Among these, he calculated for São Tomé that 25 species (64.1%) were single island endemics, with 19 (59.4%) for Príncipe; an additional 5 and 6 species respectively for each island were regarded as endemic to the Gulf of Guinea islands as a whole (i.e. present on more than one of the islands of São Tomé, Príncipe, Bioko or Annobon). GASCOIGNE (1993, 1996) also published useful bibliographies of the fauna of the islands. Most of the land-snail species were discovered and named during the nineteenth century, in publications by RANG (1831), MORELET (1848, 1858, 1868), DOHRN (1866a, 1866b), GREEFF (1882), NOBRE (1886, 1891, 1894) and GIRARD (1893a, 1893b), with reviews by CROSSE (1868, 1888a, 1888b) and Girard. In the early twentieth century, GERMAIN (1908, 1912a, 1912b, 1915, 1916) reported on additional large collections of specimens, but the proportion of new species he found was lower, so one might gain the impression that faunal exploration had almost been completed. After this there was a long break in activity, until GASCOIGNE (1994a) became resident on São Tomé and reported new discoveries of a few species that had been introduced to the islands. However, the paper by GERMAIN (1916) was the last to give a comprehensive critical checklist of the land Mollusca of either of the islands. DTH and GAH visited both islands in December 2013 and made small collections of land Mollusca, and subsequently prepared a review of the endemic genus Rhysotina of São Tomé (HOLYOAK & HOLYOAK, 2016). RFL and MP made several longer visits to the islands over the next few years, partly to study Archachatina, during which small numbers of other molluscs were obtained. Unidentified taxa from the collections made in 2013 appeared to represent undescribed species, so DTH and GAH returned to both islands in November- December 2018 in order to collect more systematically. Their fieldwork on Príncipe was carried out with assistance from FS and his co-workers from the Fundacão Príncipe and these continued to collect molluscs in other areas during 2019. RFL and MP also continued to work on both islands in 2018 and 2019, resulting in more specimens found during studies focussed on Archachatina. This paper describes the 13 new species discovered during all of this fieldwork, along with other material representing new distributional records for either island. We found that very small snails (<3 mm) had often been overlooked by earlier workers, doubtless because they mainly occur at very low densities, sieving techniques were not used in the past and the weak daylight reaching the ground inside the rainforests makes direct searching unproductive. However, some taxa with larger shells had also been overlooked, possibly because they resembled known species. Research on the material collected is continuing in order to clarify and update the taxonomy of many of the endemic species from anatomical and other characters, a process made more difficult by the poor level of taxonomic understanding of the faunas of neighbouring continental countries. Because nothing better 4 exists to fill the need for a full checklist, we give provisional full species lists for both islands in Table 1. MATERIAL AND METHODS Fieldwork was carried out by DTH and GAH on São Tomé from 1-7 and 11-13 Dec. 2013 and 24 Nov.-2 Dec. and 8-12 Dec. 2018; on Príncipe on 9-10 Dec. 2013 and 3-6 Dec. 2018. In 2018 we were assisted in the field on some days by RFL, MP and FS, along with local guides and other helpers, and these authors also made small collections independently. Fieldwork sites were selected by DTH and GAH to represent a range of molluscan habitats, but with a preference for native forest in good condition. At each site we attempted to find and record a full list of all species of non-marine Mollusca present, including slugs, and to always collect samples of those that could not be reliably identified in the field.
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